The earliest musk deer of the genus <Emphasis Type="Italic">Moschus</Emphasis> and their significance in clarifying of evolution and relationships of the family Moschidae

Moschus grandaevus Schlosser, the most ancient musk deer, is recorded from two Late Miocene localities in the south of Eastern Siberia, Olkhon Island (Lake Baikal) and Taralyk-Cher near Kyzyl (Tuva). The morphological study of the species elucidates the origin, evolution, and relationships of the genus Moschus and the entire family Moschidae. A new classification of the Moschidae is proposed.     

Who killed Micromeryx flourensianus? A case study of taphonomy and predation on ruminants in the middle Miocene of France

The fossil remains of the moschid Micromeryx flourensianus from the Sansan type locality (middle Miocene, France) show indications of fluvial transport and clear signs of carnivore activity. Most remarkable are bite marks with a distinctive pattern on the heels of the moschids (marks predominantly on the medial side of the astragalus and on the lateral side of the calcaneus). These allow the reconstruction of a specific hunting behaviour: most likely, the predator was attacking the moschid prey by deliberate bites on the heel during the chase, probably to immobilize it. This kind of behaviour is recorded for mustelids hunting moschids in modern ecosystems. Considering this and evaluating the size and pattern of the bite marks, it was possible to name the mustelids Martes sansaniensis, Taxodon sansaniensis and Proputorius sansaniensis as the most likely candidates for the hunt on Micromeryx. This offers an insight into a predator–prey relationship in the fossil‐rich locality Sansan and reveals that a hunting behaviour present in modern mustelids might actually represent a trait at least 14 million years old.     

Micromorphological,geochemical, and diagenetic characterization of sirenian ribs preserved in the Late Miocene paleontological site of Cessaniti (southern Calabria,Italy)

The site of Cessaniti (Vibo Valentia, Italy) has been well known since the 19th century for the richness and good preservation of its Miocene fauna and flora. The sedimentary succession of the site represents a paralic system that evolved toward an open-marine environment recording the Tortonian transgression. The fossil assemblage contains rich invertebrate (corals, bivalves, gastropods, brachiopods, echinoids, benthic and planktonic foraminifers) and vertebrate faunas (proboscideans, rhinoceroses, giraffids, bovids, sirenids, marine turtles, and fish remains). The fossils recovered at the Cessaniti site have a relevant role in phylogenetic studies and paleogeographic reconstructions of Late Miocene environments of the southern Italy. This research is focused on the microstructure and preservation state of the fossil bones. Samples of Metaxytherium sp. bones have been analyzed to understand the diagenetic profile of the bone assemblages that characterizes the taphonomic history of the Cessaniti site. The analyses provided a comprehensive account of how bone mineral (bioapatite) has been altered and demonstrated that the post-burial processes did not significantly affect the micromorphological and biogeochemical features of the bones. The excellent preservation state of the bones strengthens the importance of the Cessaniti site for studies of the Mediterranean Miocene vertebrate fauna.     

Paleoenvironment of Dryopithecus brancoi at Rudabánya, Hungary: evidence from dental meso- and micro-wear analyses of large vegetarian mammals

The environment of the hominoid Dryopithecus brancoi at Rudabánya (Late Miocene of Hungary) is reconstructed here using the dietary traits of fossil ruminants and equids. Two independent approaches, dental micro- and meso-wear analyses, are applied to a sample of 73 specimens representing three ruminants: Miotragocerus sp. (Bovidae), Lucentia aff. pierensis (Cervidae), Micromeryx flourensianus (Moschidae), and one equid, Hippotherium intrans (Equidae). The combination of meso- and micro-wear signatures provides both long- and short-term dietary signals, and through comparisons with extant species, the feeding styles of the fossil species are reconstructed. Both approaches categorize the cervid as an intermediate feeder engaged in both browsing and grazing. The bovid Miotragocerus sp. is depicted as a traditional browser. Although the dental meso-wear pattern of the moschid has affinities with intermediate feeders, its dental micro-wear pattern also indicates significant intake of fruits and seeds. Hippotherium intrans was not a grazer and its dental micro-wear pattern significantly differs from that of living browsers, which may suggest that the fossil equid was engaged both in grazing and browsing. However, the lack of extant equids which are pure browsers prevents any definitive judgment on the feeding habits of Hippotherium. Based on these dietary findings, the Rudabánya paleoenvironment is reconstructed as a dense forest. The presence of two intermediate feeders indicates some clearings within this forest; however the absence of grazers suggests that these clearings were most likely confined. To demonstrate the ecological diversity among the late Miocene hominoids in Europe, the diet and habitat of Dryopithecus brancoi and Ouranopithecus macedoniensis (Greece) are compared.     

First record of late middle Miocene Moschidae from Turkey: Micromeryx and Hispanomeryx from Catakbağyaka (Muğla,SW Turkey)

Here, we describe remains of Micromeryx sp. and Hispanomeryx sp. from the middle Miocene locality Catakbağyaka (MN 7/8). This is the first record for Miocene Moschidae for the late middle Miocene in Turkey and verifies the importance of Turkey as a corridor for the dispersal of Eurasian mammals. Furthermore, the record from Catakbağyaka confirms that the sympatric co-occurrence of two Miocene moschid taxa in one locality appears to be a common phenomenon.     

Leftover prey remains: a new taphonomic mode from the Late Miocene Cerro Azul Formation of Central Argentina

The Cerro Azul Formation (La Pampa Province, Argentina) comprises a rich vertebrate fauna of small mammals dominated by notoungulates and rodents. The fauna pertains to the Late Miocene specifically to the Huayquerian Stage/Age. Taphonomic analysis of micro‐mammals from Estancia Ré locality evidenced that the faunal assemblage was accumulated by the activity of a predator. This assemblage was compared with others from the Cerro Azul Formation in Telén and Caleufú localities (La Pampa Province), previously interpreted as products of predator activities. These microfossil accumulations differ from assemblages attributed to pellets and faeces produced by modern predators (nocturnal and diurnal bird raptors and carnivore mammals). However, due to their anatomical representation, degree of bone breakage and presence of tooth marks, they are interpreted as accumulations of uneaten prey remains discarded by the predator. The predator involved could not be determined with certainty, although the presence of tooth marks in some skeletal remains and the presence of coprolites in Telén and Caleufú suggest that it could be a carnivore mammal. Similarities in the accumulation mechanism, patterns of preservation and sedimentary contexts in the three assemblages support the recognition of a new taphonomic mode, termed ‘leftover prey remains’.     

Molecular and morphological phylogenies of ruminantia and the alternative position of the moschidae

The ruminants constitute the largest group of ungulates, with >190 species, and its distribution is widespread throughout all continents except Australia and Antarctica. Six families are traditionally recognized within the suborder Ruminantia: Antilocapridae (pronghorns), Bovidae (cattle, sheep, and antelopes), Cervidae (deer), Giraffidae (giraffes and okapis), Moschidae (musk deer), and Tragulidae (chevrotains). The interrelationships of the families have been an area of controversy among morphology, palaeontology, and molecular studies, and almost all possible evolutionary scenarios have been proposed in the literature. We analyzed a large DNA data set (5,322 nucleotides) for 23 species including both mitochondrial (cytochrome b, 12S ribosomal RNA (rRNA), and 16S rRNA) and nuclear (kappa-casein, cytochrome P-450, lactoferrin, and alpha-lactalbumin) markers. Our results show that the family Tragulidae occupies a basal position with respect to all other ruminant families, confirming the traditional view that separates Tragulina and Pecora. Within the pecorans, Antilocapridae and Giraffidae emerge first, and the families Bovidae, Moschidae, and Cervidae are allied, with the unexpected placement of Moschus close to bovids rather than to cervids. We used these molecular results to assess the homoplastic evolution of morphological characters within the Ruminantia. A Bayesian relaxed molecular clock approach based on the continuous autocorrelation of evolutionary rates along branches was applied to estimate the divergence ages between the major clades of ruminants. The evolutionary radiation of Pecora occurred at the Early/Late Oligocene transition, and Pecoran families diversified and dispersed rapidly during the Early and Middle Miocene. We propose a biogeographic scenario to explain the extraordinary expansion of this group during the Cenozoic era.     

Scontrone (central Italy), signs of a 9‐million‐year‐old tragedy

The fossilierous bonebeds of Scontrone (Abruzzo region, central Italy) are preserved in tidal‐flat aeolian calcarenites at the base of the Lithothamnion Limestone, a Miocene carbonate ramp widespread in the central‐southern Apennines. The site bears evidence of a catastrophic event at 9 Ma. Reported are the results of the palaeobiological and taphonomic analysis conducted on the rich vertebrate assemblage, particularly on the remains of Hoplitomeryx (Mammalia, Artiodactyla, Ruminantia), recovered from the so‐called Scontrone calcarenites between 1992 and 2012. This is the first taphonomic study of a Late Miocene continental bone assemblage preserved in coastal deposits. The bones are not in primary context. They were likely exhumed during the initial phase of a marine transgression after a period of primary ‘storage’ within a possibly flood‐generated deposit in an estuarine environment. The mortality patterns indicate that the carcasses accumulated in a short time (within a year). The bones of the disarticulated skeletons were then removed, broken in a dry and brittle state, scattered over wide carbonate ramps along an arid to semi‐arid, wind‐exposed coastline and eventually buried again in aeolian calcarenites that drape transgressive tidal‐flat creek deposits. The analysis also reveals that hoplitomerycids were possibly seasonal reproducers and that the land they inhabited, the so‐called Apulia Platform, was probably swept by sudden, disastrous, storm‐supplied flash floods.     

Functional anatomy of the cervical region in the late Miocene amphicyonid Magericyon anceps (Carnivora,Amphicyonidae): implications for its feeding behaviour

We describe the skull and neck morphology of the late Miocene amphicyonid Magericyon anceps, focusing on aspects related to functional anatomy. This species, recorded only from the Vallesian sites of Batallones‐1 and Batallones‐3 (Madrid, Spain), is the last known amphicyonid in the fossil record of Western Europe, with the Batallones populations being one of the best‐known of the family. The morphology of its skull and cervical vertebrae allows us to infer aspects of its associated musculature, such as muscle strength and range of movement. Magericyon anceps had well‐developed neck muscles, suited for providing the head with a high capacity for lateral and rotatory movements, as well as for playing an important role in the extension and stabilization of the head and neck, improving its efficiency in killing and consuming prey. Magericyon anceps shared its habitat with other large carnivorans, which would have strongly influenced its behaviour. Rapid killing and processing of prey would have been an advantage for avoiding kleptoparasitism by other large predators, as well as reducing consumption time, during which M. anceps would have been more vulnerable to attack from competitors.     

Rôle du gypaète barbu (Gypaetus barbatus) dans la constitution de l’assemblage osseux de la grotte du Noisetier (Fréchet-Aure,Hautes-Pyrénées,France)

Located in the Aure Valley (French central Pyrénées), the Noisetier cave opens at 825 m asl. This Middle Palaeolithic site has produced a Mousterian industry associated with a rich faunal assemblage. Bone remains are dominated by Chamois then Ibex and Red deer. Taphonomic analysis strongly suggests a human origin for the Ibex and the Red deer but questions arise for the Chamois. The chamois assemblage differed from the bones of both species by its low cut marks ratio. Semi-digested bones are particularly abundant. Differences are even more perceptible for skeletal portions. For Red deer and Ibex, long bone remains are mostly shaft portions whereas for Chamois, ends are the most abundant. Comparisons of the Noisetier cave bone assemblage with different present-day reference data leads to the conclusion that Chamois bones have mostly been accumulated by bearded vultures.     

The petrosal bone and bony labyrinth of early to middle Miocene European deer (Mammalia,Cervidae) reveal their phylogeny

Deer (Cervidae) have a long evolutionary history dating back to the Early Miocene, around 19 million years ago. The best known fossils to document this history belong to European taxa, which all bear cranial appendages more or less similar to today's deer antlers. Despite the good fossil record, relationships of the earliest stem deer and earliest crown deer are much debated. This hampers precise calibration against the independent evidence of the fossil record in molecular clock analyses. While much has been written on the Early and Middle Miocene deer, only two phylogenetic analyses have been performed on these taxa to date mostly based on cranial appendage characters. Because the petrosal bone and bony labyrinth have been shown to be relevant for phylogeny in ruminants, we describe for the first time these elements for four iconic early cervids from Europe (Procervulus dichotomus, Heteroprox larteti, Dicrocerus elegans and Euprox furcatus) and include them in a phylogenetic analysis based on the ear region exclusively. The analysis recovered E. furcatus in a sister position to the living red deer (Cervus elaphus). Further, it placed D. elegans in a sister position to Euprox + Cervus and a clade Procervulinae that includes P. dichotomus and H. larteti, in sister position to all other deer. The inclusion of E. furcatus in crown Cervidae, which was previously suggested based on antler morphology, cannot be ruled out here but needs a more comprehensive comparison to other crown deer to be confirmed. J. Morphol. 277:1329–1338, 2016. © 2016 Wiley Periodicals, Inc.     

Species Delimitation in the Genus Moschus (Ruminantia: Moschidae) and Its High-Plateau Origin

The authenticity of controversial species is a significant challenge for systematic biologists. Moschidae is a small family of musk deer in the Artiodactyla, composing only one genus, Moschus. Historically, the number of species in the Moschidae family has been debated. Presently, most musk deer species were restricted in the Tibetan Plateau and surrounding/adjacent areas, which implied that the evolution of Moschus might have been punctuated by the uplift of the Tibetan Plateau. In this study, we aimed to determine the evolutionary history and delimit the species in Moschus by exploring the complete mitochondrial genome (mtDNA) and other mitochondrial gene. Our study demonstrated that six species, M. leucogaster, M. fuscus, M. moschiferus, M. berezovskii, M. chrysogaster and M. anhuiensis, were authentic species in the genus Moschus. Phylogenetic analysis and molecular dating showed that the ancestor of the present Moschidae originates from Tibetan Plateau which suggested that the evolution of Moschus was prompted by the most intense orogenic movement of the Tibetan Plateau during the Pliocene age, and alternating glacial-interglacial geological eras.     

Frasnian vertebrate taphonomy and sedimentology of macrofossil concentrations from the Langsēde Cliff,Latvia

Luk?evi?s, E., Ahlberg, P.E., Stinkulis, ?., Vasi?kova, J. & Zupi??, I. 2011: Frasnian vertebrate taphonomy and sedimentology of macrofossil concentrations from the Langsēde Cliff, Latvia. Lethaia, Vol. 45, pp. 356–370. The siliciclastic sequence of the Upper Devonian of Kurzeme, Western Latvia, is renowned for abundant vertebrate fossils, including the stem tetrapods Obruchevichthys gracilis and Ventastega curonica. During the first detailed taphonomic study of the vertebrate assemblage from the Ogre Formation cropping out at the Langsēde Cliff, Imula River, abundant vertebrate remains have been examined and identified as belonging to one psammosteid, two acanthodian and three sarcopterygian genera; the placoderm Bothriolepis maxima dominates the assemblage. Besides fully disarticulated placoderm and psammosteid plates, separate sarcopterygian scales and teeth, and acanthodian spines, partly articulated specimens including complete distal segments of Bothriolepis pectoral fins, Bothriolepis head shields and sarcopterygian lower jaws have been found. The size distribution of the placoderm bones demonstrates that the individuals within the assemblage are of approximately uniform age. Distinct zones have been traced within the horizontal distribution of the bones. These linear zones are almost perpendicular to the dominant dip azimuth of the cross‐beds and ripple‐laminae and most probably correspond to the depressions between subaqueous dunes. Concavity ratio varies significantly within the excavation area. The degree of fragmentation of the bones and disarticulation of the skeletons suggest that the carcasses were reworked and slightly transported before burial. Sedimentological data suggest deposition in a shallow marine environment under the influence of rapid currents. The fossiliferous bed consists of a basal bone conglomerate covered by a cross‐stratified sandstone with mud drapes, which is in turn overlain by ripple laminated sandstone, indicating the bones were buried by the gradual infilling of a tidal channel. All the Middle–Upper Devonian vertebrate bone‐beds from Latvia are associated with sandy to clayey deposits and have been formed in a sea‐coastal zone during rapid sedimentation episodes, but differ in fossil abundance and degree of preservation. □Agnathans, Devonian, facies analysis, fish, fossil assemblage, palaeoenvironment.     

Vertebrate taphonomy in loess-palaeosol deposits: A case study from the late Miocene of central Argentina

This paper deals with the taphonomic analysis of the late Miocene bone assemblage from the Cerro Azul Formation at Telén (La Pampa Province, Argentina). The faunal assemblage was assigned to the Huayquerian mammal age (late Miocene). The fossiliferous section shows a homogeneous lithology, and is interpreted as a loess deposit with two similar and slightly developed palaeosols, classified as calcic vertisols. The studied sample comprises 5598 remains anatomically and taxonomically determined. They were collected from an area of about 48,000 m², appearing randomly distributed through the section and with low density. Most remains are small- to very small-sized, disarticulated, and very fragmented. Different taphonomical histories are inferred for microvertebrates and macromammals. The microvertebrate assemblage is interpreted as the result of predator activities. After a brief period of pre-burial exposure, remains were dispersed from the original depositional area. On the other hand, a natural and gradual death process is envisaged for macromammals, followed by a long period of exposure to weathering and dispersal by physical agents. Remains of both groups, once buried, suffered the diagenetic processes of the host rock. Consequently, the fossil assemblage from Telén would represent a condensed assemblage corresponding to two distinct time spans, i.e., the accumulation of microvertebrates took place in a short time interval whereas that of macromammals occurred over a longer period, coincident with the development of both soils.     

Ecological correlates of Himalayan musk deer Moschus leucogaster

Himalayan musk deer (Moschus leucogaster; hereafter musk deer) are endangered as a result of poaching and habitat loss. The species is nocturnal, crepuscular, and elusive, making direct observation of habitat use and behavior difficult. However, musk deer establish and repeatedly use the same latrines for defecation. To quantify musk deer habitat correlates, we used observational spatial data based on presence–absence of musk deer latrines, as well as a range of fine spatial‐scale ecological covariates. To determine presence–absence of musk deer, we exhaustively searched randomly selected forest trails using a 20‐m belt transect in different study sites within the Neshyang Valley in the Annapurna Conservation Area. In a subsequent way, study sites were classified as habitat or nonhabitat for musk deer. A total of 252 plots, 20 × 20 m, were systematically established every 100 m along 51 transects (each ~0.5 km long) laid out at different elevations to record a range of ecological habitat variables. We used mixed‐effect models and principal component analysis to characterize relationships between deer presence–absence data and habitat variables. We confirmed musk deer use latrines in forests located at higher elevations (3,200–4,200 m) throughout multiple seasons and years. Himalayan birch (Betula utilis) dominated forest, mixed Himalayan fir (Abies spectabilis), and birch forest were preferred over pure Himalayan fir and blue pine (Pinus wallichiana) forest. Greater crown cover and shrub diversity were associated with the presence of musk deer whereas tree height, diameter, and diversity were weakly correlated. Topographical attributes including aspect, elevation, distance to water source, and slope were also discriminated by musk deer. Over‐ and understory forest management can be used to protect forests likely to have musk deer as predicted by the models to ensure long‐term conservation of this rare deer.     

The fossil turtles of Greece: An overview of taxonomy and distribution

Turtle remains are common in the Miocene-Holocene deposits of Greece, and are a key focus of the growing research interest in Neogene herpetofaunas from the Aegean region. Some of the most important finds include one of Europe's stratigraphically youngest pleurodiran taxa, Nostimochelone lampra, from the Early Miocene of Macedonia, together with arguably the richest record of fossil tortoises from the Eastern Mediterranean. This incorporates the presently oldest definitive representatives of the quintessential genus Testudo sensu stricto from the Late Miocene of Attica and Macedonia, and numerous specimens of the colossal (carapace ∼2 m-length) testudinid Cheirogaster from Late Miocene-Late Pliocene sediments in southern and northern Greece, as well as on the eastern Aegean islands of Samos and Lesvos. Tantalising, but as yet unconfirmed Miocene accounts of the geoemydid Mauremys in Macedonia, and indeterminate emydid-like remains from Euboea, also provide potentially significant range extensions. Although hampered by a historically sparse documentation, the fossil turtles of Greece are a significant resource that record both assemblage changes and the origin of modern lineages.     

Lower Pleistocene stratigraphy of the Burdur Basin of SW Anatolia

The Burdur Basin is one of the NE-trending extensional orogen-top basins of SW Anatolia hosting alluvial-fan, fluvial and lacustrine deposits from the Late Miocene onward. The remains of Equus (Allohippus) sp., Paracamelus cf. gigas, and medium-sized deer (Cervidae gen. indet.) from the uppermost reach of the basin-fill succession show an interregional palaeobiogeographical linkage. The composition of fossil associations is typical for the Villafranchian of eastern Europe and central Asia. The architecture of the basin-fill stratigraphy and associated fossil taxa refines the Early Pleistocene regional palaeogeography and biodiversity of Anatolia drawn over the Eurasian migration pattern.     

Hippotraginae (Bovidae,Artiodactyla, Mammalia) from the Late Miocene of Tuva

Fossil remains of Late Miocene (Baode, NMU10-NMU11) horse antelopes from Tuva (Russia) are described, including Tragoreas sp., Protoryx tuvaensis sp. nov. from the Taralyk Cher locality and Quirliqnoria sp. from the Kholu locality. A new species, Protoryx tuvaensis Dmitrieva et Serdyuk, is described. These taxa compose a Late Miocene antelope assemblage of a new eastern geographical point (Russia, Tuva, Baode, NMU10-MN11).     

Sexual Dimorphism and Mortality Bias in a Small Miocene North American Rhino, <Emphasis Type="Italic">Menoceras arikarense</Emphasis>: Insights into the Coevolution of Sexual Dimorphism and Sociality in Rhinos

Rhinos are the only modern perissodactyls that possess cranial weapons similar to the horns, antlers and ossicones of modern ruminants. Yet, unlike ruminants, there is no clear relationship between sexual dimorphism and sociality. It is possible to extend the study of the coevolution of sociality and sexual dimorphism into extinct rhinos by examining the demographic patterns in large fossil assemblages. An assemblage of the North American early Miocene (∼22 million years ago) rhino, Menoceras arikarense, from Agate Springs National Monument, Nebraska, exhibits dimorphism in incisor size and nasal bone size, but there is no detectible dimorphism in body size. The degree of dimorphism of the nasal horn is greater than the degree of sexual dimorphism of any living rhino and more like that of modern horned ruminants. The greater degree of sexual dimorphism in Menoceras horns may relate to its relatively small body size and suggests that the horn had a more sex-specific function. It could be hypothesized that Menoceras evolved a more gregarious type of sociality in which a fewer number of males were capable of monopolizing a larger number of females. Demographic patterns in the Menoceras assemblage indicate that males suffered from a localized risk of elevated mortality at an age equivalent to the years of early adulthood. This mortality pattern is typical of living rhinos and indicates that young males were susceptible to the aggressive behaviors of dominant individuals in areas conducive to fossilization (e.g., ponds, lakes, rivers). Menoceras mortality patterns do not suggest a type of sociality different from modern rhinos although a group forming type of sociality remains possible. Among both living and extinct rhinos, the severity of socially mediated mortality seems unrelated to the degree of sexual dimorphism. Thus, sexual dimorphism in rhinos is not consistent with traditional theories about the co-evolution of sexual dimorphism and sociality.